Branching coral as a macroalgal refuge in a marginal coral reef system

Marginal coral reef systems may provide valuable insights into the nature of ecosystem processes in systems on the trajectory towards a phase shift to an alternate ecosystem state. This study investigates the process of herbivory in a marginal coral reef system in the Keppel Islands at the southern end of the Great Barrier Reef. Branching Acropora coral and the brown macroalga Lobophora variegata occupied up to 95% of the reef crest substratum at the three surveyed reefs. Feeding rates of herbivorous fishes and removal rates of Lobophora were directly quantified within areas of branching Acropora and on planar surfaces. Feeding rates by herbivorous fishes were habitat dependent with the highest bite rates being found in planar habitats for both Lobophora and the epilithic algal matrix (EAM) by 1–2 orders of magnitude, respectively. Feeding rates on Lobophora were, however, much lower than rates on the EAM. The low rates of Lobophora removal and significantly lower rates of herbivory in branching habitats were consistent with the high biomass of this brown alga throughout the Keppel Islands and with its distribution on reef crests, where Lobophora biomass was 20 times greater in branching than in planar habitats. This lack of feeding by herbivorous fishes within branching coral habitats in the Keppel Islands contrasts with the typical role of coral and topographic complexity on herbivores on coral reefs and highlights the potential for complex interactions between algae, corals and fishes on coral reefs. On marginal systems, herbivory may modify algal distributions but may be unable to contain the proliferation of algae such as Lobophora.     

Grazer biomass correlates more strongly with production than with biomass of algal turfs on a coral reef

The biomass of large herbivorous grazing fish on the shallow reef crest of Myrmidon Reef, Great Barrier Reef, is 7.0 times that on the reef slope (15 m depth), and 2.3 times that on the reef flat. Biomass of algal turfs on the crest was only 1.4 and 1.0 times that on the slope and flat, respectively. In contrast, rate of production of algal turfs on the crest was 5.3 and 2.8 times that on the slope and flat, respectively. A multiple correlation between large grazer biomass, algal turf biomass, and algal turf production across the three zones showed that only rate of algal production correlated significantly with large grazer biomass (algal production p=0.007, algal biomass p=0.418). This result suggests that large grazers may aggregate in zones of highest algal turf production. The mechanisms by which fish respond to habitat-specific differences in food production remain unclear.     

Spatial variation in the effects of grazing on epilithic algal turfs on the Great Barrier Reef, Australia

Of all benthic components on tropical reefs, algal turfs are the most widespread and the main source of primary productivity. We compared the importance of grazing by herbivores on algal turfs on two zones with marked differences in terms of benthic composition, herbivore biomass and grazing pressure, the inner flat and crest, of an inshore reef on the Great Barrier Reef, Australia. A combination of herbivore exclusion cages and transplants of coral rubble covered by algal turfs between reef zones was used to examine changes in algal turfs over a 4-day experimental period. In situ crest turfs had lower algal height, sediment loads and particulate content than reef flat turfs. Caged samples on the crest exhibited an increase in all three variables. In contrast, in situ and caged treatments on the flat presented algal turfs with similar values for the three analysed variables, with high algal height and heavy particulate and sediment loads. In the absence of cages, reef flat turfs transplanted to the crest had decreased algal height, total particulate material and particulate inorganic content, while the opposite was found in crest turf samples transplanted to the flat. Our results highlight the dynamic nature of algal turfs and the clear differences in the relative importance of herbivory in shaping turf length and sediment load between the reef crest and inner flat.     

Parrotfish predation on massive Porites on the Great Barrier Reef

Parrotfish grazing scars on coral colonies were quantified across four reef zones at Lizard Island, Northern Great Barrier Reef (GBR). The abundance of parrotfish grazing scars was highest on reef flat and crest, with massive Porites spp. colonies having more parrotfish grazing scars than all other coral species combined. Massive Porites was the only coral type positively selected for grazing by parrotfishes in all four reef zones. The density of parrotfish grazing scars on massive Porites spp., and the rate of new scar formation, was highest on the reef crest and flat, reflecting the lower massive Porites cover and higher parrotfish abundance in these habitats. Overall, it appears that parrotfish predation pressure on corals could affect the abundance of preferred coral species, especially massive Porites spp, across the reef gradient. Parrotfish predation on corals may have a more important role on the GBR reefs than previously thought.     

Direct and indirect effects of nursery habitats on coral‐reef fish assemblages,grazing pressure and benthic dynamics

Migrating species are common within seascapes, but the potential for these movements to alter the populations and functional roles of non‐migrating species (e.g. by increasing predation) is rarely investigated. This study considers whether the presence of nursery habitats (mangroves and seagrass) simply enhances the abundance of nursery‐using parrotfishes and piscivores on nearby coral reefs, or also affects other parrotfishes. Data from 131 reef sites and multiple seascape configurations across 13 degrees of latitude were used to model correlations between biophysical variables, including nursery habitat connectivity, and the abundance and grazing pressure of both nursery‐using species and other parrotfishes and piscivore biomass. Connectivity to mangroves and dense seagrass was positively correlated with the biomass of nursery‐using species, but was also negatively correlated with non‐nursery parrotfish populations. This reduction may be caused indirectly by nursery habitats increasing confamilial competition and predation by nursery‐using piscivores, particularly affecting small parrotfishes settling directly onto reefs. As key reef grazers, parrotfishes affect coral demographics. Consequently, a spatial simulation model predicted the impacts after five years of changes in grazing pressure because of nursery habitat connectivity. The model demonstrated that high nursery connectivity was correlated to changes in grazing pressure on nearby reefs that could potentially lead to differences in coral cover of ～3–4% when compared to low connectivity reefs. However, the direction of this change depended on the seascapes’ characteristics. Historically, large‐bodied, nursery‐using parrotfish would have increased grazing in all nursery‐rich seascapes. Overfishing means that nursery availability may have spatially variable impacts on coral cover, influencing reserve design. This study suggests that nursery availability may directly and indirectly modify an ecological process, and alter an ecological cascade (migrating species increase predator and competitor abundances, affecting other grazers and consequently corals). Therefore, elucidating the multi‐species impacts of animal movements is required to better understand ecosystem functioning.     

Coral reef mesopredators switch prey,shortening food chains,in response to habitat degradation

Diet specificity is likely to be the key predictor of a predator's vulnerability to changing habitat and prey conditions. Understanding the degree to which predatory coral reef fishes adjust or maintain prey choice, in response to declines in coral cover and changes in prey availability, is critical for predicting how they may respond to reef habitat degradation. Here, we use stable isotope analyses to characterize the trophic structure of predator–prey interactions on coral reefs of the Keppel Island Group on the southern Great Barrier Reef, Australia. These reefs, previously typified by exceptionally high coral cover, have recently lost much of their coral cover due to coral bleaching and frequent inundation by sediment‐laden, freshwater flood plumes associated with increased rainfall patterns. Long‐term monitoring of these reefs demonstrates that, as coral cover declined, there has been a decrease in prey biomass, and a shift in dominant prey species from pelagic plankton‐feeding damselfishes to territorial benthic algal‐feeding damselfishes, resulting in differences in the principal carbon pathways in the food web. Using isotopes, we tested whether this changing prey availability could be detected in the diet of a mesopredator (coral grouper, Plectropomus maculatus). The δ¹³C signature in grouper tissue in the Keppel Islands shifted from a more pelagic to a more benthic signal, demonstrating a change in carbon sources aligning with the change in prey availability due to habitat degradation. Grouper with a more benthic carbon signature were also feeding at a lower trophic level, indicating a shortening in food chains. Further, we found a decline in the coral grouper population accompanying a decrease in total available prey biomass. Thus, while the ability to adapt diets could ameliorate the short‐term impacts of habitat degradation on mesopredators, long‐term effects may negatively impact mesopredator populations and alter the trophic structure of coral reef food webs.     

Effects of physical and biological disturbances on algal turfs in Kaneohe Bay, Hawaii

Disturbance in coral reef environments commonly results in an algal community dominated by highly productive, small filamentous forms and cyanobacteria, collectively known as algal turf. Research on the types of disturbance responsible for this community structure has concentrated mainly on biological disturbance in the form of grazing, although physical and other forms of biological disturbances may be important in many coral reef areas. On the reef flat in Kaneohe Bay, Oahu, Hawaii, algal turfs grow primarily upon coral rubble that tumbles with passing swells. We manipulated the frequency of rubble tumbling in field experiments to mimic the effects of physical disturbance by abrasion and light reduction on algal biomass, canopy height, and community structure. Treatments approximated a gradient of disturbance intensities and durations that occur on the reef flat. Although sea urchins and herbivorous fishes are not widespread and abundant on the reef flat, biological disturbances to algal turf communities in the form of herbivory by small crabs and abrasion by tough macroalgae contributed significantly to the variation in algal turf biomass. Within all experiments increasing disturbance significantly reduced algal biomass and canopy heights and the community structure shifted to more disturbance-tolerant algal forms. This study shows that the chronic physical disturbances from water motion and biological disturbances other than grazing from large herbivores can control algal communities in coral reef environments.     

Territorial behavior of threespot damselfish (Eupomacentrus planifrons) increases reef algal biomass and productivity

Synopsis Algal growth and damselfish (Eupomacentrus planifrons) territories were studied in two reef habitats at Discovery Bay, Jamaica. Damselfish territories were contiguous in the reef flat (0 to 2.5 m), where the algal composition and biomass varied from territory to territory. In contrast, on the lower reef terrace (22 m), damselfish territories were often spatially segregated. While the algal composition of the territories was more uniform on the reef terrace, the total algal biomass was lower than in the territories on the reef flat. Damselfish are largely herbivorous, and they defend their territories against most intruding fish, including a number of herbivorous species. Areas of the reef terrace outside of damselfish territories were heavily grazed by herbivorous fishes and contained only small quantities of non-crustose algae.The reef terrace territories were characterized by a multispecific turf of algae (greens, blue-greens, and reds) covering the Acropora cervicornis framework and by the leafy, brown alga, Lobophora variegata. A rapid reduction in the biomass of brown algae and filamentous algae was noted when damselfish were permanently removed from their territories. Only calcified, encrusting algae — plants apparently somewhat undesirable as fish food sources — would be common on the terrace zone of this reef if damselfish territories were absent. Damselfish territoriality may significantly influence the dynamics of some reefs by increasing the biomass of the algal turf thereby increasing; reef productivity. Since blue-green algae, potential nitrogen fixers, occur in these algal turfs, the fish may also be indirectly affecting reef nutrition.     

Trophic significance of herbivorous macroinvertebrates on the central Great Barrier Reef

The common herbivorous macroinvertebrates on reef flats of the central Great Barrier Reef (GBR) were, in order of abundance: gastropod molluscs (Turbo and Trochus spp.); pagurid crabs; and the starfish, Linckia laevigata. The density of macroinvertebrates on Davies Reef was lowest in the thick-turf habitats at the windward reef-crest (0.6–0.9 m^–2) compared with 3.1 to 5.2 m^–2 elsewhere on the reef flat. Invertebrate grazer densities were similar on mid-shelf reef flats (mean: 2.3–3.6 m^–2) and significantly lower on outer-shelf reefs (0.3–1.0 m^–2). The ingestion rate of Turbo chrysostomus, the most abundant macroinvertebrate species, was derived from (a) faecal production and food absorption efficiency, (b) comparison of algal biomass on grazed and ungrazed natural substrata and (c) gut-filling rate and feeding periodicity in field populations. The ingestion rate of Trochus pyramis, the most common trochid and an abundant component of the macroinvertebrate fauna, was also estimated using (a). This gastropod fed continuously, whereas T. chrysostomus showed a distinct nocturnal feeding periodicity. T. chrysostomus and T. pyramis ingested daily means of 35 and 54 mgC animal^-1, respectively. Total gastropod grazing rates (mgC m^–2d^-1 in the field ranged from 11 in a thick-turf, reef-crest habitat to 144 on the open-grazed main flat. Grazing by gastropods accounted for between 0.3 and 8% of the net production of benthic algal food resources, depending on location on the reef flat. Across the whole reef flat the mean (areally-weighted) gastropod grazing rate was 6% of net production. A comparison of the relative roles of different types of grazers led to the conclusion that fishes are likely to have the greatest overall trophic impact on reefs of the central GBR. Even where macroinvertebrates are most abundant on reef flats, the yield from benthic algal communities to macroinvertebrates is estimated to be only one third of that due to fishes.Contribution no. 471 from the Australian Institute of Marine Science     

Meiofauna sediment relations in leeward slope turf algae of Heron Island reef

As part of studies investigating the influence of grazers on reef meiofauna, we assessed the density, composition and richness of meiofauna (retained on a 100-μm sieve) on the leeward reef slope of Heron Reef, GBR, Australia using an airlift vacuum sampling device. Estimates of meiofauna densities ranged between 40 individuals 10 cm⁻² and 290 individuals 10 cm⁻², which is considerably lower than many estimates from carbonate sediments and hard substrates from other reefs and marine habitats. The 17 taxa of meiofauna were dominated by harpacticoid copepods (40%) and nematodes (32%). Varying sediment load within algal turfs explained 37% of variation of meiofauna density. A model is proposed in which increased shelter afforded by high living coral cover reduces meiofaunal losses from grazing and increases sediment loads, balanced by areas of low coral cover in which sedimentation rates are lower and grazing rates higher. At none of the four sites did major differences in abundance occur between November and March sampling events. Together these observations suggest that epilithic meiofaunal communities are generally spatially and temporally predictable at small scales in this reef system, indicating that their ecological services are similarly conservative. Handling editor: I. Nagelkerken     

Depth-dependent indicators of algal turf herbivory throughout the Main Hawaiian Islands

Herbivorous fish are key to maintaining a balance between coral and algae on reefs, where reefs with greater herbivore biomass often show lower algal cover. For reefs worldwide, algal turf cover is expanding and is increasingly used as an indicator of disturbance. Water depth affects reef fish composition; thus, it may be expected that herbivory could also differ by depth. We examined relationships between algal turf cover and biomass (g m⁻²), density (# m⁻²) and size (cm) of herbivore groups (grazers, browsers and scrapers) across shallow (< 6 m), mid (6–18 m) and deep (18–30 m) coral reefs in the Main Hawaiian Islands. We find that across all depth classes, algal turf cover decreased with increasing grazer and scraper density, with steeper relationships observed at mid and deep reefs than in shallow reefs. In contrast, algal turf cover slightly increased with increasing grazer and browser biomass at deep reefs. Considering fish size, algal turf cover increased with larger grazer and scrapers at mid and deep reefs. The results indicate that herbivorous fish density, rather than biomass, is a better indicator of reductions in algal turf cover and resulting coral-algal balance on Hawaiian reefs, where smaller fish exert greater top-down control on cover than larger fish. Despite significant differences in herbivorous fish compositions, length-frequency distributions and fishing intensities across depth, algal turf cover remains similar across depths. Increases in fishing would have a disproportionately negative impact in deep than shallow reefs due to a lower overall fish density, where grazing functions in deep reefs are maintained by significantly fewer and smaller grazers and browsers, and larger scrapers, than in shallow reefs. Developing an understanding of patterns of algal turf herbivory by depth is important to understanding the spatial scale at which herbivory and regime shifts operate.

     

Direct evaluation of macroalgal removal by herbivorous coral reef fishes

Few studies have examined the relative functional impacts of individual herbivorous fish species on coral reef ecosystem processes in the Indo-Pacific. This study assessed the potential grazing impact of individual species within an inshore herbivorous reef fish assemblage on the central Great Barrier Reef (GBR), by determining which fish species were able to remove particular macroalgal species. Transplanted multiple-choice algal assays and remote stationary underwater digital video cameras were used to quantify the impact of local herbivorous reef fish species on 12 species of macroalgae. Macroalgal removal by the fishes was rapid. Within 3 h of exposure to herbivorous reef fishes there was significant evidence of intense grazing. After 12 h of exposure, 10 of the 12 macroalgal species had decreased to less than 15% of their original mass. Chlorodesmis fastigiata (Chlorophyta) and Galaxaura sp. (Rhodophyta) showed significantly less susceptibility to herbivorous reef fish grazing than all other macroalgae, even after 24 h exposure. Six herbivorous and/or nominally herbivorous reef fish species were identified as the dominant grazers of macroalgae: Siganus doliatus, Siganus canaliculatus, Chlorurus microrhinos, Hipposcarus longiceps, Scarus rivulatus and Pomacanthus sexstriatus. The siganid S. doliatus fed heavily on Hypnea sp., while S. canaliculatus fed intensively on Sargassum sp. Variation in macroalgal susceptibility was not clearly correlated with morphological and/or chemical defenses that have been previously suggested as deterrents against herbivory. Nevertheless, the results stress the potential importance of individual herbivorous reef fish species in removing macroalgae from coral reefs.     

Size-structural shifts reveal intensity of exploitation in coral reef fisheries

Fisheries exploitation represents a considerable threat to coral reef fish resources because even modest levels of extraction can alter ecological dynamics via shifts of stock size, species composition, and size-structure of the fish assemblage. Although species occupying higher trophic groups are known to suffer the majority of exploitative effects, changes in composition among lower trophic groups may be major, though are not frequently explored. Using size-based biomass spectrum analysis, we investigate the effects of fishing on the size-structure of coral reef fish assemblages spanning four geopolitical regions and determine if patterns of exploitation vary across trophic groups. Our analyses reveal striking evidence for the variety of effects fisheries exploitation can have on coral reef fish assemblages. When examining biomass spectra across the entire fish assemblage we found consistent evidence of size-specific exploitation, in which large-bodied individuals experience disproportionate reductions. The pattern was paralleled by and likely driven by, strongly size-specific reductions among top predators. In contrast, evidence of exploitation patterns was variable among lower trophic groups, in many cases including evidence of reductions across all size classes. The breadth of size classes and trophic groups that showed evidence of exploitation related positively to local human population density and diversity of fishing methods employed. Our findings highlight the complexity of coral reef fisheries and that the effects of exploitation on coral reefs can be realized throughout the entire fish assemblage, across multiple trophic groups and not solely restricted to large-bodied top-predators. Size-specific changes among fishes of lower trophic groups likely lead to altered ecological functioning of heavily exploited coral reefs. Together these findings reinforce the value of taking a multi-trophic group approach to monitoring and managing coral reef fisheries.     

Are the oral teeth of combtooth bennies (Salariini: Teleostei) merely combs?

Careful analysis of gut contents has resulted in the reclassification of several species of grazing fish as detritivores, shifting the trophodynamic assignment of many prominent reef grazers. Combtoothed blennies, which are among the most numerous grazing fish of the Great Barrier Reef, have been shown to target the detrital component of the epilithic algal matrix (EAM). It has been suggested that blennies have specialized dental morphology that allows them to comb through fronds of algae, collecting detritus, while leaving the algal component intact. In this study, we analysed the capability of a common reef flat blenny, Salarias fasciatus, to remove algae by (i) examining oral morphology for evidence of wear and adaptations for abrasion, and (ii) a short-term EAM feeding experiment. Examination of S. fasciatus teeth with scanning electron microscopy (SEM) showed evidence of macrowear (changes in tooth height or shape), microwear (surface chips and striae), and tooth replacement that suggests abrasion on the substrate. Energy Dispersive X-ray Spectrometry (EDS) revealed that oral teeth incorporate mineral iron. When fed artificial substrata with a developed EAM layer S. fasciatus removed 57% of photosynthetic material and 38.5% of organic biomass. Although studies have found that blenny gut contents consist predominantly of detritus, blennies are still likely to contribute to the removal of algae on coral reefs.     

Habitat associations of juvenile fish at Ningaloo Reef, Western Australia: the importance of coral and algae

Habitat specificity plays a pivotal role in forming community patterns in coral reef fishes, yet considerable uncertainty remains as to the extent of this selectivity, particularly among newly settled recruits. Here we quantified habitat specificity of juvenile coral reef fish at three ecological levels; algal meadows vs. coral reefs, live vs. dead coral and among different coral morphologies. In total, 6979 individuals from 11 families and 56 species were censused along Ningaloo Reef, Western Australia. Juvenile fishes exhibited divergence in habitat use and specialization among species and at all study scales. Despite the close proximity of coral reef and algal meadows (10's of metres) 25 species were unique to coral reef habitats, and seven to algal meadows. Of the seven unique to algal meadows, several species are known to occupy coral reef habitat as adults, suggesting possible ontogenetic shifts in habitat use. Selectivity between live and dead coral was found to be species-specific. In particular, juvenile scarids were found predominantly on the skeletons of dead coral whereas many damsel and butterfly fishes were closely associated with live coral habitat. Among the coral dependent species, coral morphology played a key role in juvenile distribution. Corymbose corals supported a disproportionate number of coral species and individuals relative to their availability, whereas less complex shapes (i.e. massive & encrusting) were rarely used by juvenile fish. Habitat specialisation by juvenile species of ecological and fisheries importance, for a variety of habitat types, argues strongly for the careful conservation and management of multiple habitat types within marine parks, and indicates that the current emphasis on planning conservation using representative habitat areas is warranted. Furthermore, the close association of many juvenile fish with corals susceptible to climate change related disturbances suggests that identifying and protecting reefs resilient to this should be a conservation priority.     

Maintenance of fish diversity on disturbed coral reefs

Habitat perturbations play a major role in shaping community structure; however, the elements of disturbance-related habitat change that affect diversity are not always apparent. This study examined the effects of habitat disturbances on species richness of coral reef fish assemblages using annual surveys of habitat and 210 fish species from 10 reefs on the Great Barrier Reef (GBR). Over a period of 11 years, major disturbances, including localised outbreaks of crown-of-thorns sea star (Acanthaster planci), severe storms or coral bleaching, resulted in coral decline of 46–96% in all the 10 reefs. Despite declines in coral cover, structural complexity of the reef framework was retained on five and species richness of coral reef fishes maintained on nine of the disturbed reefs. Extensive loss of coral resulted in localised declines of highly specialised coral-dependent species, but this loss of diversity was more than compensated for by increases in the number of species that feed on the epilithic algal matrix (EAM). A unimodal relationship between areal coral cover and species richness indicated species richness was greatest at approximately 20% coral cover declining by 3–4 species (6–8% of average richness) at higher and lower coral cover. Results revealed that declines in coral cover on reefs may have limited short-term impact on the diversity of coral reef fishes, though there may be fundamental changes in the community structure of fishes.     

Coral–macroalgal phase shifts or reef resilience: links with diversity and functional roles of herbivorous fishes on the Great Barrier Reef

Changes from coral to macroalgal dominance following disturbances to corals symbolize the global degradation of coral reefs. The development of effective conservation measures depends on understanding the causes of such phase shifts. The prevailing view that coral–macroalgal phase shifts commonly occur due to insufficient grazing by fishes is based on correlation with overfishing and inferences from models and small-scale experiments rather than on long-term quantitative field studies of fish communities at affected and resilient sites. Consequently, the specific characteristics of herbivorous fish communities that most promote reef resilience under natural conditions are not known, though this information is critical for identifying vulnerable ecosystems. In this study, 11 years of field surveys recorded the development of the most persistent coral–macroalgal phase shift (>7 years) yet observed on Australia’s Great Barrier Reef (GBR). This shift followed extensive coral mortality caused by thermal stress (coral bleaching) and damaging storms. Comparisons with two similar reefs that suffered similar disturbances but recovered relatively rapidly demonstrated that the phase shift occurred despite high abundances of one herbivore functional group (scraping/excavating parrotfishes: Labridae). However, the shift was strongly associated with low fish herbivore diversity and low abundances of algal browsers (predominantly Siganidae) and grazers/detritivores (Acanthuridae), suggesting that one or more of these factors underpin reef resilience and so deserve particular protection. Herbivorous fishes are not harvested on the GBR, and the phase shift was not enhanced by unusually high nutrient levels. This shows that unexploited populations of herbivorous fishes cannot ensure reef resilience even under benign conditions and suggests that reefs could lose resilience under relatively low fishing pressure. Predictions of more severe and widespread coral mortality due to global climate change emphasize the need for more effective identification and protection of ecosystem components that are critical for the prevention of coral reef phase shifts.     

Enhancement of Grazing Gastropod Populations as a Coral Reef Restoration Tool: Predation Effects and Related Applied Implications

We aimed to evaluate the efficacy of the gastropod grazer Trochus niloticus in controlling epilithic algae and enhancing coral recruitment on artificial substrata on coral reefs where the biomass of herbivorous fishes was low due to heavy fishing pressure. Hatchery‐reared, subadult trochus were stocked onto pallet balls (small artificial reefs composed of concrete and limestone aggregate) at a density of approximately four individuals per square meter (external surface area). This density was re‐established with releases of new trochus each month for 6 months. At the end of the experiment, there were no significant differences in algal biomass, cover and community composition, or the density of coral recruits on substrata with and without trochus. High monthly attrition of stocked trochus on the pallet balls, apparently due mainly to predation by octopus, did not allow the evaluation of the efficiency of the trochus enhancement, at the desired density, as a restoration tool. However, at the lower trochus densities (circa 1 m^?2), which occurred as a result of predation in this study, no apparent enhancement of algal grazing or coral recruitment were observed. The surprisingly high predation of stocked trochus in a heavily fished and gleaned reef site stresses the importance of understanding all the factors affecting the survival of stocked animals. To help mitigate predation of trochus, artificial habitat with refuge spaces that allow the grazers to escape predation could be provided and individuals of a larger size could be released.     

Non‐reef habitats in a tropical seascape affect density and biomass of fishes on coral reefs

Nonreef habitats such as mangroves, seagrass, and macroalgal beds are important for foraging, spawning, and as nursery habitat for some coral reef fishes. The spatial configuration of nonreef habitats adjacent to coral reefs can therefore have a substantial influence on the distribution and composition of reef fish. We investigate how different habitats in a tropical seascape in the Philippines influence the presence, density, and biomass of coral reef fishes to understand the relative importance of different habitats across various spatial scales. A detailed seascape map generated from satellite imagery was combined with field surveys of fish and benthic habitat on coral reefs. We then compared the relative importance of local reef (within coral reef) and adjacent habitat (habitats in the surrounding seascape) variables for coral reef fishes. Overall, adjacent habitat variables were as important as local reef variables in explaining reef fish density and biomass, despite being fewer in number in final models. For adult and juvenile wrasses (Labridae), and juveniles of some parrotfish taxa (Chlorurus), adjacent habitat was more important in explaining fish density and biomass. Notably, wrasses were positively influenced by the amount of sand and macroalgae in the adjacent seascape. Adjacent habitat metrics with the highest relative importance were sand (positive), macroalgae (positive), and mangrove habitats (negative), and fish responses to these metrics were consistent across fish groups evaluated. The 500‐m spatial scale was selected most often in models for seascape variables. Local coral reef variables with the greatest importance were percent cover of live coral (positive), sand (negative), and macroalgae (mixed). Incorporating spatial metrics that describe the surrounding seascape will capture more holistic patterns of fish–habitat relationships on reefs. This is important in regions where protection of reef fish habitat is an integral part of fisheries management but where protection of nonreef habitats is often overlooked.